Monday, October 15, 2018

[Herpetology • 2018] Multilocus Phylogeny and Revised Classification for Mountain Dragons of the Genus Japalura s.l. (Agamidae: Draconinae) from Asia

Japalura s.l. 

in Wang, Che, Lin, Deepak, Aniruddha, et al., 2018. 

Although the genus Japalura s.l. has long been recognized as paraphyletic based on limited genetic sampling, its problematic taxonomy has not been revised, and phylogenetic relationships among the majority of congeners remain unknown. Here we utilize a densely sampled dataset of both multilocus genetic and morphological data to provide the first phylogenetic inference of relationships among Japalura s.l.species. Our results show that Japalura s.l. is paraphyletic, consisting of four major clades that are scattered across the phylogeny of the subfamily Draconinae: the first clade from the western, central and middle-eastern Trans-Himalayas, the second clade from the far eastern Trans-Himalayas, the third clade from East Asia and the last clade from Indochina. To address this widespread paraphyly of the genus and to stabilize the taxonomy within the family Draconinae, we revise the current taxonomy and split Japalura s.l. into four genera. By doing so, we recognize two existing generic names, Japalura sensu stricto and Pseudocalotes, resurrect one name available in the literature, Diploderma, and describe one new genus, Cristidorsa gen. nov. We discuss phylogenetic relationships and taxonomy within Japalura s.l. and present a diagnostic key to all recognized genera of the subfamily Draconinae.

Keywords: China, India, integrative taxonomy, lizard, new genus, Tibetan Plateau

Figure 2. Phylogenetic relationships among Japalura sensu lato based on both Maximum Likelihood and Bayesian Analyses of two mitochondrial genes (COI and ND2), tRNAs and three nuclear genes (BDNF, CMOS, and R35). Maximum Likelihood bootstrap and Bayesian posterior probability values are included at all nodes, except (1) terminal nodes that unify multiple individuals of the same species [which all have 1.00/100 support (Bayesian/Maximum Likelihood)]; and (2) non-conflicting (either kind of analysis yielded significant supports) yet inconsistent nodes between two types of analyses (e.g. nodes unifying clades I and J), in which only the Bayesian posterior probability is given. Photos of selected species of Japalura s.l. are included (scaled to reflect relative sizes of each species) with their current taxonomic names (vs. revised taxonomic names, summarized in taxonomic accounts).

Taxonomic accounts 
Japalura Gray, 1853
 Etymology: The Latin name ‘Japalura’ may be derived from a locality name in India, and the term is feminine gender. We suggest the English common name as ‘Himalayan Dragon’, and the Chinese name as ‘攀蜥’ (pronounced as ‘Pan-Xi’). 

Type species: Japalura variegata, Gray, 1853.

Included species: Based on our phylogenetic results, we assign the following species to the genus Japalura sensu stricto: J. andersoniana, J. kumaonensis, J. tricarinata and J. variegata. Following our morphological results and proposed morphological diagnoses, we also assign J. dasi, J. major and J. sagittifera into this genus, pending future phylogenetic studies.

Cristidorsa Wang, Deepak, Datta-Roy, Lin, Jiang, Che & Siler gen. nov.

 Etymology: The Latin term ‘Cristidorsa’ means ‘ridged dorsum’, which describes the distinct, characteristic ridges on the dorsal surface of the body in the new genus. The generic name is feminine and it consists of two parts, namely ‘Cristi-’ (meaning ‘ridged’) and ‘-dorsa’ (meaning ‘dorsum’). We suggest the English common name as ‘Ridged Dragons’ and the Chinese name as ‘棱背蜥’ (pronounced as ‘Leng-Bei-Shi’). 

Type species: Cristidorsa otai (Mahony, 2009).

Included species: Based on our phylogenetic results, we assign C. otai and C. planidorsata to the genus Cristidorsa.

Diploderma Hallowell, 1861 

Etymology: The Latin generic name ‘Diploderma’ consists of two parts, ‘Diplo-’ means ‘double’ or ‘many’, and ‘-derma’ means ‘skin’, and the whole word is in a neuter gender. As the previous generic name ‘Japalura’ and most species names of the genus s.l. are feminine, most names of species that are now assigned to Diploderma need their gender changed to neutral (except for existing neutral-gender names like brevipes or flaviceps, Latin nouns like vela or names derived from peoples’ names, i.e. dymondi, luei, makii, swinhonis, varcoae and zhaoermii). We suggest the English common name of the genus as ‘Mountain Dragon’, and the Chinese common name as ‘龙蜥’ (pronounced as ‘Long-Xi’). 

Type species: Diploderma polygonatum Hallowell, 1861.

Included species: Based on our phylogenetic results, we assign the following species into the genus Diploderma: D. batangense, D. brevipes, D. chapaense, D. dymondi, D. flaviceps, D. laeviventre, D. luei, D. makii, D. micangshanense, D. polygonatum (and all of its subspecies), D. slowinskii, D. splendidum, D. swinhonis, D. varcoae, D. vela, D. yulongense, D. yunnanense and D. zhaoermii

According to our proposed morphological diagnoses, we also assign Diploderma brevicaudum, D. fasciatum, D. grahami, D. hamptoni and D. iadinum to this genus, pending on future phylogenetic confirmations. In total, 24 species of Japalura s.l. are reclassified into the genus Diploderma (for discussion on the taxonomic status of D. ngoclinense see below).

Pseudocalotes Fitzinger, 1843 

Type species: Pseudocalotes tympanistriga (Gray, 1831).

Included species: A single species of Japalura s.l. Pseudocalotes kingdonwardi bapoensis, is reclassified into the genus Pseudocalotes. Currently, the genus includes 22 recognized species in total (Grismer et al., 2016a; Harvey et al., 2017).

Kai Wang, Jing Che, Simin Lin, V. Deepak, Datta-Roy Aniruddha, Ke Jiang, Jieqiong Jin, Hongman Chen and Cameron D. Siler. 2018. Multilocus Phylogeny and Revised Classification for Mountain Dragons of the Genus Japalura s.l. (Reptilia: Agamidae: Draconinae) from Asia. Zoological Journal of the Linnean Society. zly034.  DOI:  10.1093/zoolinnean/zly034 

[Botany • 2018] Thismia kelabitiana (Thismiaceae) • A New Unique Fairy Lantern from Borneo Potentially Threatened by Commercial Logging

Thismia kelabitiana Dančák, Hroneš & Sochor

in Dančák, Hroneš, Sochor & Sochorová, 2018.
Pa’Umor Fairy Lantern  ||  DOI:  10.1371/journal.pone.0203443 

Thismia kelabitiana, a new unique species from the Sarawak state of Malaysia in the island of Borneo is described and illustrated. This new species is not similar to any species of Thismia described so far especially by having a unique form of mitre and outer perianth lobes deeply divided into 8–10 acute lobes and forming striking fringe around perianth tube opening. The species appears to be critically endangered due to ongoing logging activities in the region. It may potentially become a surrogate species for lower montane forests of the region and thus help protect them against further destruction.

Fig 1. Thismia kelabitiana. A, Plant with flower bud. B, Plant with young flower. C, Plant with mature flower. D, Whole plant with root system. E, Detail of mitre and perianth opening. Photos Michal Sochor. 

Fig 3. Thismia kelabitiana. A, Habit of flowering plant. B, Side view of flower. C, Inner view of stamens. D, Style with stigma. E, Outer view of stamens. Drawn by Kateřina Janošíková.

Fig 2. Thismia kelabitiana. A, Outer view of stamens showing the lateral appendages and apical parts of connectives. B, Young capsule with persistent stigma. C, Inner view of stamens showing ribbed connectives. D, Seeds inside mature capsule. Photos Michal Sochor. 


Thismia kelabitiana Dančák, Hroneš & Sochor, sp. nov. 

Type. MALAYSIA, Sarawak: Kelabit Highlands, Pa'Umor village, Anak Kadi Ridge, 4.4 km SSE of the village. Elevation 1195 m a.s.l., 13 January 2017. M. Sochor, M. Hroneš, M. Dančák, Z. Egertová & J.R. Pasan BOR1/17 (holotype SAR [in spirit and herbarium specimen, accession number Sochor/BOR-1/17], isotype OL [35272]).

Diagnosis. Thismia kelabitiana differs significantly from all congeneric species by the combination of the following traits, e.g., flowers large (up to 2.8 × 1.8 cm), outer perianth lobes deeply divided into 8–10 acute lobes and forming striking fringe around perianth tube opening, mitre relatively small and flat elevated by three long filiform pillars, connectives with prominent longitudinal rib and three appendices on apical margin.

Fig 4. Habitat of Thismia kelabitiana. Ravine of a small stream in lower montane tropical rain forest. Photo Michal Sochor.

Habitat and ecology. The species occurs in lower montane primary tropical rainforest at an altitude around 1200 m a. s. l. It was found in humid stream ravines as well as in relatively drier open forest sites (Fig 4). A variety of other mycoheterotrophic species were abundant at the type locality, including Aphyllorchis pallida, Burmannia lutescens agg., B. championii, Cystorchis aphylla, Gymnosiphon aphyllus agg., Epirixanthes kinabaluensis, Exacum tenue, Platanthera saprophytica, Sciaphila arfakiana, S. cf. nana, S. tenella, Thismia cornuta, T. minutissima ined., T. aff. nigra and T. viridistriata. Herbaceous vegetation was otherwise sparse. Thismia kelabitiana seems not to prefer any particular environmental conditions at the locality as it occurs in various aspects of slopes with various inclinations either in rugged ravines or relatively flat terrain in various distances from a stream.

Distribution. The species is known only from the type locality and the other, 600 m distant locality, where plants have only been photographed. Both localities are found on a ridge south-east from Pa’Umor village in the Bario district of Sarawak (Malaysia).

Etymology. The specific epithet reflects the geographical origin of the species in the Kelabit Highlands, the land of the Kelabit people.

Common name. Pa’Umor Fairy Lantern. There is no widely accepted generic common name for the genus Thismia. The recent wave of interest as well as potential conservational employment, nevertheless, calls for this name. Sometimes the name Fairy Lantern is used [Thiele & Jordan, 2012, Lưu et al., 2014] which has originally belonged to Thismia rodwayi (either in singular or plural form as Fairy Lanterns). Since the name has already been used for a few other Thismia species, we follow this approach. We suggest calling Thismia kelabitiana Pa’Umor Fairy Lantern. The name is derived from Pa’Umor village community in whose forest the species occurs.

Martin Dančák, Michal Hroneš, Michal Sochor and Zuzana Sochorová. 2018. Thismia kelabitiana (Thismiaceae), A New Unique Fairy Lantern from Borneo Potentially Threatened by Commercial Logging.  PLoS ONE. 13(10): e0203443.  DOI:  10.1371/journal.pone.0203443

[Botany • 2018] Hsuehochloa calcarea (Poaceae, Bambusoideae, Arundinarieae) • A New Genus of Temperate Woody Bamboos from A Limestone Montane Area of China

Hsuehochloa calcarea (C. D. Chu & C. S. Chao) D. Z. Li & Y. X. Zhang

in Zhang, Ma & Li, 2018.

Ampelocalamus calcareus is a climbing and slender bamboo, known from south Guizhou, China. This species grows in broadleaved forests of limestone montane areas. Recent molecular phylogenetic analyses demonstrated that A. calcareus was sister to all other lineages of the tribe Arundinarieae rather than a member of Ampelocalamus. The morphological features and habitats of A. calcareus and related genera including Ampelocalamus, Drepanostachyum and Himalayacalamus were compared and discussed. The characteristics of the branch complements, nodes and foliage leaves distinguish A. calcareus from morphologically similar taxa. On the basis of molecular and morphological evidence, we propose to establish a new genus, Hsuehochloa, to accommodate A. calcareus and to honour the late Chinese bamboo taxonomist Chi-Ju Hsueh (Ji-Ru Xue). In addition, we describe the inflorescence of Hsuehochloa for the first time.

Keywords: Ampelocalamus, climbing bamboos, Hsuehochloa, new genus

Figure 1. Hsuehochloa calcarea.
A, B Habit and habitat C Clump D Young culm with white pubescence E, F Branch complement G Culm sheath H Leaves I Inflorescence J Floret
(A–D, G from P. F. Ma & Z. M. Cai 10050 E, F, H from seedlings introduced from Libo, Guizhou, China I, J from P. F. Ma s.n). Scale bars: 5 cm (A–C); 0.5 cm (D, G); 2 cm (E, F); 1 cm (H, I); 1mm (J).


Hsuehochloa D. Z. Li & Y. X. Zhang, gen. nov.

Diagnosis: Hsuehochloa resembles genera Ampelocalamus, Drepanostachyum and Himalayacalamus, but differs from those genera by its thin culms (4–5 mm), fewer branches in each branch complement (1, 3–7), inconspicuous nodal sheath scar, falcate auricles and leathery foliage leaves.

Type: Hsuehochloa calcarea (C. D. Chu & C. S. Chao) D. Z. Li & Y. X. Zhang, comb. nov.

Basionym. Ampelocalamus calcareus C. D. Chu & C. S. Chao, 
1983 Acta Phytotax. Sin. 21: 204–206. 
Type: CHINA, Guizhou, Libo, 500 m, C. D. Chu, C. S. Chao, J. Q. Zhang & K. M. Lan 81018 (holotype, NF!; isotype, PE!)
Etymology: Hsuehochloa was named in honour of the late Prof. Chi-Ju Hsueh (Ji-Ru Xue in Pinyin transliteration) (1921–1999), a pioneer Chinese botanist on bamboos of SW China and mentor of the senior author in 1983–1986. Hsueh stands for his family name and chloa means grass.

Distribution and habitat: Endemic to south Guizhou, China, under broadleaved forests in a limestone montane area at 500–950 m altitude.

 Yu-Xiao Zhang, Peng-Fei Ma and De-Zhu Li. 2018. A New Genus of Temperate Woody Bamboos (Poaceae, Bambusoideae, Arundinarieae) from A Limestone Montane Area of China. PhytoKeys. 109: 67-76.  DOI: 10.3897/phytokeys.109.27566

Friday, October 12, 2018

[Paleontology • 2018] The Smallest Diplodocid Skull Reveals Cranial Ontogeny and Growth-Related Dietary Changes in the Largest Dinosaurs

Life reconstruction of CMC VP14128. Note the cranial morphologies interpreted to denote differing feeding strategies: in CMC VP14128 the narrow snout with posteriorly elongated and morphologically varied tooth row for bulk feeding vs. the widened snout with anteriorly restricted peg-shaped teeth for ground-level browsing in adults. Also note the camouflaged ontogenetic color change suggesting young diplodocids may have sought forested refuge.

in Woodruff, Carr, Storrs, et al., 2018. 
Reconstruction by A. Atuchin.

Sauropod dinosaurs were the largest terrestrial vertebrates; yet despite a robust global fossil record, the paucity of cranial remains complicates attempts to understand their paleobiology. An assemblage of small diplodocid sauropods from the Upper Jurassic Morrison Formation of Montana, USA, has produced the smallest diplodocid skull yet discovered. The ~24 cm long skull is referred to cf. Diplodocus based on the presence of several cranial and vertebral characters. This specimen enhances known features of early diplodocid ontogeny including a short snout with narrow-crowned teeth limited to the anterior portion of the jaws and more spatulate teeth posteriorly. The combination of size plus basal and derived character expression seen here further emphasizes caution when naming new taxa displaying the same, as these may be indicative of immaturity. This young diplodocid reveals that cranial modifications occurred throughout growth, providing evidence for ontogenetic dietary partitioning and recapitulation of ancestral morphologies.

Figure 1: Skeletal reconstruction of CMC VP14128 to scale with a mature Diplodocus carnegii (dark grey). Grey bones are missing, while those in ivory are those present in CMC VP14128. Skeletal reconstruction based on the Diplodocus by S. Hartman. Silhouettes by S. Hartman and PhyloPic (Creative Commons Attribution-ShareAlike 3.0 Unported;, modifications made. Skeletal reconstruction of CMC VP14128 redrawn from D. carnegii skeletal by S. Hartman ( Human scale is Andrew Carnegie at his natural height of 1.6 m. Skeletal and silhouettes to scale. (B) CMC VP14128 in right lateral view with accompanying schematic. (C) CMC VP14128 in left lateral view with accompanying schematic. Schematics by DCW. The four portions of the skull numbered on accompanying schematics.

 Lateral views and schematics to scale. a: angular, al: alisphenoid, aof: antorbital fenestra, d: dentary, f: frontal, h: hyoid, l: lacrimal, m: maxilla, n: nasal, oc: occipital condyle, os: orbitosphenoid, p: parietal, paof: preantorbital fenestra, pf: prefrontal, pm: premaxilla, po: postorbital, pro: prootic, q: quadrate, sa: surangular, sq: squamosal. L and r before bone denotes if it is left or right.

Systematic Paleontology

Saurischia Seeley 1887
Sauropodomorpha von Huene 1932

Sauropoda Marsh 1878
Diplodocoidea Marsh 1884
Flagellicaudata Harris and Dodson 2004

Diplodocidae Marsh 1884

cf. Diplodocus Marsh 1878.


Figure 5: Life reconstruction of CMC VP14128. Note the cranial morphologies interpreted to denote differing feeding strategies: in CMC VP14128 the narrow snout with posteriorly elongated and morphologically varied tooth row for bulk feeding vs. the widened snout with anteriorly restricted peg-shaped teeth for ground-level browsing in adults. Also note the camouflaged ontogenetic color change suggesting young diplodocids may have sought forested refuge. Reconstruction by A. Atuchin.

Within Dinosauria, there are small bodied taxa that display basal and derived characters and occupy unusual basal phylogenetic positions. The validity and position of such taxa has been disputed, and regarding sauropodomorph phylogeny, we would advocate that the combination of basal and derived characters and basal phylogenetic recovery should be recognized as an indicator of an immature ontogimorph – instead of a distinct taxon. In light of the current wealth of information pertaining to dinosaur ontogeny, we can no longer assume that all morphological differences correspond with phylogenetic distinctiveness. Accounting for ontogeny could prove as test for our phylogenies. Recognizing the ontogenetic age of immature specimens provides important insights into the life history of these animals. The immature Diplodocus specimen CMC VP14128 extends our understanding of the ontogeny of the genus and the evolution of diplodocids into new areas, where:

(1) The combination of basal and derived characters in the juvenile is broadly congruent with the phylogenetic transition from eusauropods to diplodocoids.
(2) The plesiomorphic tooth morphology is retained in immature Diplodocus and lost with maturity, and we predict this growth pattern will be seen in all other diplodocoids.
(3) As first proposed by Whitlock et al. (2010), tooth and skull morphology indicate that during growth Diplodocus inhabited different trophic levels/niches, where juveniles were generalists (i.e., browsers; Fig. 5) and more mature individuals were specialists (i.e., ground-level browsing), a pattern that we predict is ancestral for Diplodocoidea.

D. Cary Woodruff, Thomas D. Carr, Glenn W. Storrs, Katja Waskow, John B. Scannella, Klara K. Nordén and John P. Wilson. 2018. The Smallest Diplodocid Skull Reveals Cranial Ontogeny and Growth-Related Dietary Changes in the Largest Dinosaurs. Scientific Reports. 8, Article number: 14341. DOI:  10.1038/s41598-018-32620-x 

Tiny Skull Illuminates the Lives of Giant Dinosaur via @SmithsonianMag


Thursday, October 11, 2018

[PaleoMammalogy • 2018] Miopetaurista neogrivensis • Oldest Skeleton of A Fossil Flying Squirrel Casts New Light on the Phylogeny of the Group

Miopetaurista neogrivensis Mein 1970

in Casanovas-Vilar, Garcia-Porta, Fortuny, et al., 2018. 

Flying squirrels are the only group of gliding mammals with a remarkable diversity and wide geographical range. However, their evolutionary story is not well known. Thus far, identification of extinct flying squirrels has been exclusively based on dental features, which, contrary to certain postcranial characters, are not unique to them. Therefore, fossils attributed to this clade may indeed belong to other squirrel groups. Here we report the oldest fossil skeleton of a flying squirrel (11.6 Ma) that displays the gliding-related diagnostic features shared by extant forms and allows for a recalibration of the divergence time between tree and flying squirrels. Our phylogenetic analyses combining morphological and molecular data generally support older dates than previous molecular estimates (~23 Ma), being congruent with the inclusion of some of the earliest fossils (~36 Ma) into this clade. They also show that flying squirrels experienced little morphological change for almost 12 million years.

Fig 1: The fossil flying squirrel Miopetaurista neogrivensis.
 (a) Reconstruction of the skeleton based in the partial skeleton IPS56468 from Abocador de Can Mata. Missing elements are based on extant giant flying squirrel Petaurista petaurista and are colored in blue.
(b) Life appearance of Miopetaurista neogrivensis showing the animal ready to land on a tree branch. Coat pattern and color are based in extant Petaurista species, the sister taxon of Miopetaurista.  Scale bar is 4 cm.

Fig 3: Mandible and cheek teeth of Miopetaurista neogrivensis.
 (a to c) Partial left hemimandible (IPS56468j) in lateral, medial and dorsal views. (d to e) Partial right hemimandible (IPS56468i) in lateral and medial views. A caudal vertebra and a bone fragment are attached to the lateral side of the mandibular ramus. Both hemimandibles were associated to the partial skeleton IPS56468 from ACM/C5-D1. (f to g) Partial hemimandible (IPS87560) from ACM/C8-B sector in lateral and medial views. (h) Left upper cheek teeth series (P3–M3) of IPS56468h (Figure 6—Figure supplement 1 ). (i) Left lower cheek teeth series (p4–m3) of IPS56468j. Cheek teeth measurements are given in Supplementary file 4 whereas mandibular measurements are given in Supplementary file 6. For a detailed description and comparisons of cheek teeth and mandible morphology see Appendix 3.1 and 3.2. an, angular process; ar, articular process; co, coronoid process. Scale bar is 1 cm in figs. a to g; 2 mm in (h to i).

Fig 7: Flying squirrel phylogeny and node dating estimates based on a Bayesian total evidence analysis including Miopetaurista neogrivensis.

Fig 8: Fossil record of ‘flying squirrels’ and paleoclimatic data. Temporal ranges of purported flying squirrel genera in Europe, Asia and North America. The 95% highest posterior density (HPD) intervals for flying squirrel divergence as derived from total evidence and node dating analyses are indicated in orange shading (see Figure 7 and Figure 7—figure supplement 1 ). Darker shading indicates the time interval where both independently calculated estimates overlap, thus defining the most likely time interval for flying squirrel divergence. Global paleoclimatic data are taken from Zachos et al., 2001.

Miopetaurista neogrivensis is the oldest unquestionable flying squirrel and dates back to the middle/late Miocene boundary (11.6 Ma). Its diagnostic wrist anatomy indicates that the two subtribes of flying squirrels had already diverged at that time. Moreover, this new fossil allows for a recalibration of flying squirrel time of origin and diversification, generally providing somewhat older estimates than previous molecular analyses. These differ according to the phylogenetic method used, total evidence analysis estimates an interval of 36.6 – 24.9 Ma while node dating results in a younger estimate of 30.6 – 17.4 Ma. Therefore, we cannot rule out that at least some of the oldest (ca. 36 Ma) fossils tentatively identified as flying squirrels may indeed belong to this group. However, the estimates of both independent phylogenetic approaches overlap for the late Oligocene (31 – 25 Ma), which should be considered the most likely interval for flying squirrel divergence. The two flying squirrel subtribes are found to have diverged during the early Miocene (22 – 18 Ma) while most extant genera would do so during the Miocene, although they are not recorded until the Pleistocene. Miopetaurista neogrivensis is estimated to have diverged from Petaurista spp., its sister taxon, between 18.8 – 12.4 Ma, the oldest boundary overlapping with the earliest record of the genus Miopetaurista (18 – 17 Ma). Perhaps not surprisingly, the skeletons of both genera show little differences. Sciurids are often regarded as a morphologically conservative group and flying squirrels are no exception having experienced few morphological changes for almost 12 million years.

Isaac Casanovas-Vilar, Joan Garcia-Porta, Josep Fortuny, Óscar Sanisidro, Jérôme Prieto, Marina Querejeta, Sergio Llácer, Josep M Robles, Federico Bernardini, and David M Alba. 2018. Oldest Skeleton of A Fossil Flying Squirrel Casts New Light on the Phylogeny of the Group.  eLife. 7; e39270 DOI:  10.7554/eLife.39270.001

Oldest fossil of a flying squirrel sheds new light on its evolutionary tree via @elife @EurekAlert

Wednesday, October 10, 2018

[Botany • 2018] Blakea echinata (Melastomataceae: Blakeeae) • A New Species from the Caribbean Rainforest of Panama

Blakea echinata  Almeda & Penneys

in Almeda & Penneys, 2018. 

Blakea echinata from the lowland Caribbean rainforest of Panama is described, illustrated, mapped, and compared with superficially similar species. It is readily distinguished by its elongate internodes; indumentum of spreading smooth (sometimes gland-tipped) trichomes on distal branchlets, leaves, floral bracts, and calyx lobes intermixed with laterally compressed and somewhat roughened conic to clavate or ± triangular trichomes on distal branchlets, floral peduncles, and abaxial surfaces of floral bracts; rhombic reflexed basally clawed petals; yellow-orange unappendaged anthers; and ovary apex that is elaborated into a distally glandular-laciniate collar 5–6 mm long that envelops the style base. A conservation assessment of Critically Endangered is recommended for this species based on IUCN Red List Categories and Criteria.

Keywords: Blakeeae, conservation, epiphyte, neotropics, new species, Panama, Eudicots

Figure 1. Blakea echinata.
A. Habit. B. Representative leaf (abaxial surface) C. Distal portion of peduncle and outer floral bracts. D. Inner floral bracts. E. Hypanthium and calyx lobes. F. Longitudinal section of flower showing ovary, collar, and calyx lobes. G. Flower at anthesis. H. Stamens, ventral view (left) and profile view (right). I. Petal (adaxial surface).
All drawn from De Gracia 765.

Figure 2. Images of Blakea echinata.
A. Flower. B. Habit. C. Flowers and subtending leaves. D. Hypanthia and subtending floral bracts. E. Flower showing reflexed petals (right) and flower at post anthesis (left). F. Young fruiting hypanthia with spreading calyx lobes and subtending floral bracts.
Image credits: A–F (De Gracia 763) by J. E. De Gracia.

Blakea echinata Almeda & Penneys, sp. nov. 

Diagnosis: Distinguished by its epiphytic habit with elongate internodes, distal branchlets moderately to sparingly covered with a mixture of smooth (sometimes gland-tipped) spreading trichomes (1–3 mm long) that are white when fresh but dull yellow-brown when dry, and early deciduous laterally compressed and somewhat roughened conic to clavate or ± triangular trichomes up to 0.5 mm long, inner and outer floral bracts that are connate basally for 2–6 mm to form a tight collar enveloping the hypanthium, calyx tube 2–3 mm long, oblong-obovate calyx lobes, rhombic commonly reflexed petals with a basal claw, yellow-orange unappendaged anthers, and ovary apex elaborated into a distally glandular-laciniate collar 5–6 mm long that envelops the style base.

Figure 3. Geographic distribution of Blakea echinata.

Habitat and distribution:— This species has been collected in mature secondary rainforest in the Caribbean lowlands of Panama at 100–150 m elevation where it is known from a small area in the Donoso District of Colón province (Figure 3). 

 Etymology:—The epithet for this species, echinata, highlights the copious and conspicuous indumentum of stiff smooth and/or gland-tipped spreading trichomes on leaves, petioles, floral bracts, and calyx lobes.

Frank Almeda and Darin Penneys. 2018. Blakea echinata (Melastomataceae: Blakeeae): A New Species from the Caribbean Rainforest of Panama.  Phytotaxa. 372(1); 104–110. DOI:  10.11646/phytotaxa.372.1.9

Resumen: Se describe e ilustra Blakea echinata de la selva baja del Caribe de Panamá, se presenta un mapa y se la compara con especies superficialmente similares. Se distingue fácilmente por sus entrenudos alargados, indumento de tricomas suaves extendidos (a veces con punta glandular) en las ramitas distales, hojas, brácteas florales y lóbulos del cáliz entremezclados con tricomas cónicos a clavados o triangulares lateralmente comprimidos y algo rugosos en ramitas distales, pedúnculos florales y superficies abaxiales de las brácteas florales; pétalos rómbicos basalmente con una reflexa, anteras de color amarillonaranja sin apéndice y el ápice de ovario que se elabora en un collar distalmente glandular-laciniado de 5–6 mm de largo que envuelve la base de estilo. Se recomienda una categoría de conservación de En Peligro Crítico para esta especie en base a las Categorías y Criterios de la Lista Roja de la UICN.


Tuesday, October 9, 2018

[Herpetology • 2018] Micryletta nigromaculata • A New Limestone-dwelling Species of Micryletta (Anura: Microhylidae) from northern Vietnam

Micryletta nigromaculata 
Poyarkov, Nguyen, Duong, Gorin & Yang, 2018

DOI: 10.7717/peerj.5771 

We report on a new species of the genus Micryletta from limestone karst areas in northern Vietnam, which is described on the basis of molecular and morphological evidence. Micryletta nigromaculata sp. nov. is restricted to narrow areas of subtropical forests covering karst massifs in Cat Ba National Park (Hai Phong Province) and Cuc Phuong National Park (Ninh Binh Province) at elevations of 90–150 m a.s.l. In the phylogenetic analyses, the new species is unambiguously positioned as a sister lineage to all remaining species of Micryletta. We also discuss genealogical relationships and taxonomic problems within the genus Micryletta, provide molecular evidence for the validity of M. erythropoda and discuss the taxonomic status of M. steinegeri. We suggest the new species should be considered as Endangered (B1ab(iii), EN) following the IUCN’s Red List categories. A discussion on herpetofaunal diversity and conservation in threatened limestone karst massifs in Southeast Asia is provided.

Figure 4: Holotype of Micryletta nigromaculata sp. nov. (ZMMU A5934), male, in situ in dorsolateral view.
Photo by Nikolay A. Poyarkov.

Figure 6: Color variation of Micryletta nigromaculata sp. nov. in life. Cat Ba National Park:
 (A) Male paratype ZMMU A5945; (B) male paratype ZMMU A5935 in situ; Cuc Phuong National Park; (C) male DTU 302 in situ; (D) female DTU 303 in situ.
Photos A–B by Nikolay A. Poyarkov; C–D by Tan Van Nguyen.

Figure 3: Holotype of Micryletta nigromaculata sp. nov. (ZMMU A5934), male, in life.
(A) Dorsal view; (B) ventral view; (C) lateral view of head; (D) volar view of left hand; (E) plantar view of right foot.
Photos by Nikolay A. Poyarkov.

Micryletta nigromaculata sp. nov.

Diagnosis. The new species is assigned to the genus Micryletta by the following combination of morphological features: small body size; vomerine teeth absent; tympanum small, rounded, externally visible; very prominent subarticular tubercles on fingers and toes; three well-developed metacarpal tubercles; distinct supernumerary palmar and metatarsal tubercles posterior to base of digits; first finger not reduced; digit tips expanded to very small disks and webbing on fingers and toes totally absent (Dubois, 1987; Fei et al., 2009). Micryletta nigromaculata sp. nov. is distinguished from all of its congeners by a combination of the following morphological characters: body size small (SVL 18.5–23.0 mm in males, 24.2–25.9 mm in females); body habitus moderately slender; head wider than long; snout obtusely rounded in profile; EL equal to or shorter than SL; IOD two times wider than UEW; tibiotarsal articulation of adpressed limb reaching the level of eye center; dorsal surface slightly granular with small round flattened tubercles; supratympanic fold present, thick, glandular; outer metatarsal tubercle absent; dorsum coloration brown to reddish-brown; dorsum with dark-brown irregular hourglass-shaped pattern edged with orange; body flanks brown with dark-brown to black patches or spots edged with white, a large black blotch in inguinal area on each side; lateral sides of head immaculate reddish brown lacking white patches; venter whitish with indistinct gray pattern; and throat in males whitish with light-gray marbling.

Figure 2: Phylogenetic BI tree of Micryletta reconstructed on the base of 947 bp of 16S rRNA gene. Values on the branches correspond to BI PP/ML BS, respectively. For specimen, locality and GenBank accession number information see Table 1.
Photos by Nikolay A. Poyarkov (Micryletta nigromaculata sp. nov. M. erythropoda, M. cf. inornata) and Chung-Wei You (M. steinegeri).

Figure 1: Distribution of the genus Micryletta and the new species.
(A) Map of Southeast Asia with approximate range of the genus Micryletta shown in red. Black circles indicate type localities of the currently recognized taxa within Micryletta. Yellow stars show distribution of Micryletta nigromaculata  sp. nov. black dot in the center of icon indicates the type locality (Cat Ba Island). Black square indicates the inset shown in detail in B.
 (B) Map of northern Vietnam, showing distribution of Micryletta nigromaculata sp. nov. and the Red River basin; 1—Cat Ba National Park, Hai Phong Province (type locality); 2—Cuc Phuong National Park, Ninh Binh Province. Photo by Nikolay A. Poyarkov.

Etymology: Specific epithet “nigromaculata” is an adjective in the nominative case, feminine gender, derived from Latin words “niger” for “black” and “maculatus” for “spotted,” in reference the characteristic black blotches on flanks in the new species.

Recommended vernacular names: We recommend “Black-spotted Paddy Frog” as the common English name of the new species and the common name in Vietnamese as “Nhái bầu hông đen.”

Distribution and biogeography: The presently known distribution of Micryletta nigromaculata sp. nov. is shown in Fig. 1. To date, the new species is known from limestone karst areas covered by primary tropical forest in Cat Ba N. P., Hai Phong Province, and by secondary tropical forest in Cuc Phuong N. P., Ninh Binh Province at elevations 90–150 m a.s.l. Northern Vietnam has one of the world largest areas of limestone landscapes, covered by specific limestone vegetation (Fenart et al., 1999; Day & Urich, 2000). The currently known range of Micryletta nigromaculata sp. nov. is divided by the vast lowlands of the Red River valley, an important biogeographic border in Indochina (Bain & Hurley, 2011; Yuan et al., 2016); our phylogenetic analysis estimates genetic divergence between the Cat Ba and Cuc Phuong populations at 0.7% (see Table 1). It is anticipated that Micryletta nigromaculata sp. nov. also occurs in the adjacent limestone karsts of northern Vietnam; in particular, records from Quang Ninh, Lang Son and Bac Giang provinces of northeastern Vietnam, as well as from Hoa Binh, Ha Nam and Thanh Hoa provinces of northwestern Vietnam are anticipated.

Natural history notes: Our knowledge on the biology of Micryletta nigromaculata sp. nov. is scarce; the species appears to be closely associated with karstic habitats. In Cat Ba N. P. (Hai Phong Province) during a 2-week survey in October 2011, specimens were only recorded from a small patch of limestone outcrops ca. 20 m in diameter, near a large limestone karst cliff and a small temporary body of water. Frogs were observed from 16:00 to 20:00 h hiding between small pieces of limestone rocks. Despite intensive search from 10 to 22 of October 2013, no additional specimens of the new species were recorded from other areas in Cat Ba N. P. In Cuc Phuong N. P. (Ninh Binh Province) specimens were found at night between 19:00 and 23:30 h near cave entrances and in valleys surrounded by limestone cliffs, relatively near to water sources. Surrounding habitat was limestone karst covered with primary polydominant tropical forest with multilayered canopy and an abundance of lianas, with occasional trees of Streblus macrophyllus (Moraceae), Terminalia myriocarpa (Combretaceae), Parashorea chinensis (Dipterocarpaceae) and Tetrameles nudiflora (Tetramelaceae) (in Cat Ba N.P.) or secondary forest (in Cuc Phuong N.P.). Reproduction biology, including advertisement call, tadpole morphology, as well as diet of the new species remains unknown.

Other species of anurans recorded syntopically with the new species at the type locality included Polypedates megacephalus Hallowell, P. mutus (Smith), Theloderma albopunctatum (Liu & Hu), Liuixalus calcarius Milto, Poyarkov, Orlov & Nguyen, Philautus catbaensis Milto, Poyarkov, Orlov & Nguyen, Hyla chinensis Günther, Microhyla butleri Boulenger, M. heymonsi Vogt and Micryletta cf. inornata. In Cuc Phuong National Park (Ninh Binh Province) the new species was recorded in sympatry with Occidozyga martensii (Peters), Leptobrachium guangxiense Fei, Mo, Ye & Jiang, Ophryophryne microstoma Boulenger, Glyphoglossus (formerly Calluella) guttulatus (Blyth), Microhyla heymonsi Vogt, Micryletta cf. inornata (Boulenger); Rana johnsi Smith; Sylvirana cf. annamitica Sheridan & Stuart; Raorchestes cf. menglaensis (Kou); Theloderma albopunctatum (Liu & Hu) and T. annae Nguyen, Pham, Nguyen, Ngo & Ziegler.

Limestone karst areas are recognized as arks of highly endangered though still insufficiently studied biodiversity. Unique geological structure of karst massifs, formed by erosion and subterranean water drainages create numerous humid microrefugia with stable environmental conditions, which serve as an important environmental buffer for small vertebrates during periods of climate change (Clements et al., 2006; Glaw, Hoegg & Vences, 2006). The complex terrain of isolated karstic hills and caves create multiple ecological niches what along with their highly fragmented habitat-island nature result in high degrees of site-specific endemism within, and diversity among them (Oliver et al., 2017; Grismer et al., 2018). Limestone karsts are also known as important “biodiversity arks” for both surface and cave faunas, yet karstic regions are rapidly becoming some of the most imperiled ecosystems on the planet (Clements et al., 2006; Grismer et al., 2016a, 2016b, 2018; Luo et al., 2016; Suwannapoom et al., 2018). South-east Asia harbors more limestone karsts than anywhere else on earth (Day & Urich, 2000) with numerous new species including relic lineages of amphibians and reptiles being discovered from limestone areas (e.g. see discussions in Milto et al., 2013; Grismer et al., 2014; Grismer & Grismer, 2017; Grismer et al., 2016a, 2016b, 2017, 2018; Nazarov et al., 2014, 2018; Connette et al., 2017; Suwannapoom et al., 2018 and references therein). Ironically, though acting as major biodiversity hotspots, limestone karsts are critically endangered due to unregulated quarrying mostly for cement manufacturing, which is the primary threat to the survival of karst-associated species (Grismer et al., 2018); their continued exploitation for limestone cannot be stopped (Clements et al., 2006). Until karst habitats in Vietnam are thoroughly investigated, a significant portion of this country’s herpetological diversity will remain underestimated and unprotected. Our study thus calls for urgent focused survey and conservation efforts on karst herpetofauna in Southeast Asia and in Vietnam in particular.

Nikolay A. Poyarkov, Tan Van Nguyen, Tang Van Duong, Vladislav A. Gorin and Jian-Huan Yang. 2018. A New Limestone-dwelling Species of Micryletta (Amphibia: Anura: Microhylidae) from northern Vietnam. PeerJ. 6:e5771.  DOI: 10.7717/peerj.5771


[Paleontology • 2018] Dynamoterror dynastes • A New Tyrannosaurid (Dinosauria: Theropoda) from the Upper Cretaceous Menefee Formation of New Mexico

Dynamoterror dynastes 
McDonald​, Wolfe & Dooley, 2018
Invictarx zephyri  
McDonald​ & Wolfe, 2018
Art by Brian Engh 

The giant tyrannosaurids were the apex predators of western North America and Asia during the close of the Cretaceous Period. Although many tyrannosaurid species are known from numerous skeletons representing multiple growth stages, the early evolution of Tyrannosauridae remains poorly known, with the well-known species temporally restricted to the middle Campanian-latest Maastrichtian (∼77–66 Ma). The recent discovery of a new tyrannosaurid, Lythronax argestes, from the Wahweap Formation of Utah provided new data on early Campanian (∼80 Ma) tyrannosaurids. Nevertheless, the early evolution of Tyrannosauridae is still largely unsampled. We report a new tyrannosaurid represented by an associated skeleton from the lower Campanian Allison Member of the Menefee Formation of New Mexico. Despite fragmentation of much of the axial and appendicular skeleton prior to discovery, the frontals, a metacarpal, and two pedal phalanges are well-preserved. The frontals exhibit an unambiguous autapomorphy and a second potential autapomorphy that distinguish this specimen from all other tyrannosaurids. Therefore, the specimen is made the holotype of the new genus and species Dynamoterror dynastes. A phylogenetic analysis places Dynamoterror dynastes in the tyrannosaurid subclade Tyrannosaurinae. Laser-scanning the frontals and creation of a composite 3-D digital model allows the frontal region of the skull roof of Dynamoterror to be reconstructed.

 Reconstructed frontal complex of Dynamoterror dynastes.
 Missing elements reconstructed based upon Teratophoneus curriei (UMNH VP 16690) (Loewen et al., 2013). 

 Dynamoterror dynastes Invictarx zephyri 
Art by Brian Engh 

Dinosauria Owen, 1842, sensu Baron, Norman & Barrett, 2017
Theropoda Marsh, 1881, sensu Baron, Norman & Barrett, 2017
Coelurosauria Huene, 1914, sensu Sereno, McAllister & Brusatte, 2005

Tyrannosauroidea Osborn, 1906,
 sensu Walker, 1964; Sereno, McAllister & Brusatte, 2005
Tyrannosauridae Osborn, 1906,
 sensu Sereno, McAllister & Brusatte, 2005
Tyrannosaurinae Osborn, 1906,
sensu Matthew & Brown, 1922; Sereno, McAllister & Brusatte, 2005

Dynamoterror dynastes gen. et sp. nov.

Holotype: UMNH VP 28348, incomplete associated skeleton including the left and right frontals, four fragmentary vertebral centra, fragments of dorsal ribs, right metacarpal II, supraacetabular crest of the right ilium, unidentifiable fragments of long bones, phalanx 2 of left pedal digit IV, and phalanx 4 of left pedal digit IV.

Etymology: Dynamoterror is derived from the transliterated Greek word dynamis (“power”) and the Latin word terror. The specific name, dynastes, is a Latin word meaning “ruler.” The intended meaning of the binomen is “powerful terror ruler.” The name also honors the binomen “Dynamosaurus imperiosus” (Osborn, 1905), a junior synonym of Tyrannosaurus rex (Osborn, 1905, 1906), but a particular childhood favorite of the lead author.

Locality: UMNH VP 28348 was collected in San Juan County, New Mexico, on land administered by the US BLM. Precise locality data are on file at UMNH and the BLM.

Horizon: UMNH VP 28348 was collected from outcrops of the Juans Lake Beds (Miller, Carey & Thompson-Rizer, 1991), upper part of the Allison Member, Menefee Formation; lower Campanian, Upper Cretaceous. Lucas et al. (2005) produced a radioisotopic date of 78.22 ± 0.26 Ma from a bentonite layer near the top of the Menefee Formation in the Gallina hogback in the eastern part of the San Juan Basin. In the part of the San Juan Basin where UMNH VP 28348 was collected, the overlying Cliff House Sandstone contains fossils of the ammonoid Baculites perplexus (Siemers & King, 1974), corresponding to between 78.0 and 78.5 Ma (Molenaar et al., 2002). According to the regional stratigraphic correlation chart of Molenaar et al. (2002), the Menefee Formation spans approximately 84.0–78.5 Ma, based upon correlations with marine biostratigraphic zones. This age range corresponds to uppermost Santonian—middle Campanian (Cohen et al., 2013).

Specific diagnosis (as for genus by monotypy): tyrannosaurine tyrannosaurid distinguished by two autapomorphies on the frontals: (1) prefrontonasal and prefrontolacrimal processes are in close proximity, separated only by a shallow notch; and (2) subrectangular, concave, laterally projecting caudal part of the postorbital suture, separated from the rostral part by a deep groove. The second autapomorphy should be treated as provisional, given the ontogenetic variation observed in this region of the frontal in other tyrannosaurids (Carr & Williamson, 2004) (see description of the lateral surface of the frontal below). In the context of the phylogenetic analysis of Carr et al. (2017), which is used herein, UMNH VP 28348 exhibits a feature that supports its affinities among derived tyrannosauroids (1561, “frontal, dorsotemporal fossa, medial extension, dorsal view: meets opposing fossa at the midline”; also present in Timurlengia euotica, Xiongguanlong baimoensis, B. sealeyi, and Tyrannosauridae), and a feature identified by Carr et al. (2017) as an unambiguous synapomorphy of “derived tyrannosaurines” (1571, “frontal, sagittal crest, form, dorsal and lateral views: present and pronounced (dorsoventrally tall), single structure”).

The description of Dynamoterror dynastes from the lower Campanian Allison Member of the Menefee Formation provides additional data on the morphology and diversity of early tyrannosaurines in Laramidia. However, additional discoveries are needed to elucidate the paleobiogeographic history of tyrannosaurines.

Andrew T. McDonald​, Douglas G. Wolfe and Alton C. Dooley Jr. 2018. A New Tyrannosaurid (Dinosauria: Theropoda) from the Upper Cretaceous Menefee Formation of New Mexico. PeerJ. 6:e5749.  DOI:  10.7717/peerj.5749
Newly Discovered Tyrant Dinosaur Stalked Ancient New Mexico via @SmithsonianMag

Andrew T. McDonald​ and Douglas G. Wolfe. 2018. A New Nodosaurid Ankylosaur (Dinosauria: Thyreophora) from the Upper Cretaceous Menefee Formation of New Mexico. PeerJ. 6:e5435.  DOI:  10.7717/peerj.5435